Testing foraging optimization models in brown bears: Time for a paradigm shift in nutritional ecology?

Abstract

How organisms obtain energy to survive and reproduce is fundamental to ecology, yet researchers use theoretical concepts represented by simplified models to estimate diet and predict community interactions. Such simplistic models can sometimes limit our understanding of ecological principles. We used a polyphagous species with a wide distribution, the brown bear (Ursus arctos), to illustrate how disparate theoretical frameworks in ecology can affect conclusions regarding ecological communities. We used stable isotope measurements (δ13C, δ15N) from hairs of individually monitored bears in Sweden and Bayesian mixing models to estimate dietary proportions of ants, moose, and three berry species to compare with other brown bear populations. We also developed three hypotheses based on predominant foraging literature, and then compared predicted diets to field estimates. Our three models assumed (1) bears forage to optimize caloric efficiency (optimum foraging model), predicting bears predominately eat berries (~70% of diet) and opportunistically feed on moose (Alces alces) and ants (Formica spp. and Camponotus spp; ~15% each); (2) bears maximize meat intake (maximizing fitness model), predicting a diet of 35%–50% moose, followed by ants (~30%), and berries (~15%); (3) bears forage to optimize macronutrient balance (macronutrient model), predicting a diet of ~22% (dry weight) or 17% metabolizable energy from proteins, with the rest made up of carbohydrates and lipids (~49% and 29% dry matter or 53% and 30% metabolizable energy, respectively). Bears primarily consumed bilberries (Vaccinium myrtillus; 50%–55%), followed by lingonberries (V. vitis‐idaea; 22%–30%), crowberries (Empetrum nigrum; 8%–15%), ants (5%–8%), and moose (3%–4%). Dry matter dietary protein was lower than predicted by the maximizing fitness model and the macronutrient balancing model, but protein made up a larger proportion of the metabolizable energy than predicted. While diets most closely resembled predictions from optimal foraging theory, none of the foraging hypotheses fully described the relationship between foraging and ecological niches in brown bears. Acknowledging and broadening models based on foraging theories is more likely to foster novel discoveries and insights into the role of polyphagous species in ecosystems and we encourage this approach.

Trophic adaptation of large terrestrial omnivores to global change

“…Despite the signal of trophic adaptation of populations to environmental change in our study, it remains unclear to what extent populations of species that live at the physiological or ecological limits of their range in extreme environments (e.g., arctic, desert, or alpine environments) are able to cope with future changes in environmental conditions. Their adaptive capacity will depend on the magnitude and velocity of environmental change, as well as on whether alternative food sources meet the nutritional requirements of consumers (4, 46). For instance, it has been suggested that the switch of polar bears from marine prey to terrestrial food sources (e.g., berries or bird nests) in response to reduced sea-ice extent due to climate change is associated with declines in body size, body condition and reproduction (47).…”